Biol Invasions
DOI 10.1007/s10530-011-9981-2
ORIGINAL PAPER
Alien flora of India: taxonomic composition, invasion status
and biogeographic affiliations
Anzar A. Khuroo • Zafar A. Reshi •
Akhtar H. Malik • Ewald Weber • Irfan Rashid
G. H. Dar
Received: 30 September 2010 / Accepted: 2 March 2011
Ó Springer Science+Business Media B.V. 2011
Abstract The wide knowledge gaps in invasion
biology research that exist in the developing world
are crucial impediments to the scientific management
and global policymaking on biological invasions. In
an effort to fill such knowledge gaps, we present here
an inventory of the alien flora of India, based on
systematic reviews and rigorous analyses of research
studies (ca. 190) published over the last 120 years
(1890–2010 AD), and updated with field records of
the last two decades. Currently, the inventory com-
prises of 1,599 species, belonging to 842 genera in
Electronic supplementary material The online version of
this article (doi:10.1007/s10530-011-9981-2) contains
supplementary material, which is available to authorized users.
A. A. Khuroo (&)  A. H. Malik  G. H. Dar
Department of Botany, Centre for Biodiversity
and Taxonomy, University of Kashmir, Srinagar 190 006,
Jammu and Kashmir, India
e-mail: anzarak@gmail.com
Z. A. Reshi
Department of Botany, Biological Invasions Laboratory,
University of Kashmir, Srinagar 190 006,
Jammu and Kashmir, India
E. Weber
Institute of Biochemistry and Biology, Biodiversity
Research, University of Potsdam, Maulbeerallee 1,
14469 Potsdam, Germany
I. Rashid
Department of Botany, Government College Baramulla,
Baramulla 193 101, Jammu and Kashmir, India
•
161 families, and constitutes 8.5% of the total Indian
vascular flora. The three most species-rich families
are Asteraceae (134 spp.), Papilionaceae (114 spp.)
and Poaceae (106 spp.), and the three largest genera
are Eucalyptus (25 spp.), Ipomoea (22 spp.), and
Senna (21 spp.). The majority of these species (812)
have no report of escaping from cultivation. Of the
remaining subset of 787 species, which have either
escaped from intentional cultivation, or spread after
unintentional introduction, casuals are represented by
57 spp., casual/naturalised by 114 spp., naturalised by
257 spp., naturalised/invasive by 134 spp., and
invasive by 225 spp. Biogeographically, more than
one-third (35%) of the alien flora in India has its
native ranges in South America, followed by Asia
(21%), Africa (20%), Europe (11%), Australia (8%),
North America (4%); and cryptogenic (1%). The
inventory is expected to serve as the scientific
baseline on plant invasions in India, with implications
for conservation of global biodiversity.
Keywords Biodiversity  Biological invasions 
Alien plants  Taxonomy  Biogeography  India
Introduction
In invasion ecology, both taxonomic and geograph-
ical knowledge gaps exist at a global scale (Pyšek
et al. 2008), and countries of the developing world
lag far behind countries of the developed world in
123

research on invasive species (Nuñez and Pauchard
2010). In the developing world, especially Asian and
African countries, except South Africa, are poorly
represented in the scientific literature on biological
invasions. Such a data deficit in the invasion biology
literature does not, however, indicate that developing
countries are at a lower risk of being invaded by
invasive alien species, because biological invasions
are a phenomenon that transcends political bound-
aries (Rashid et al. 2009). Rather, the dearth of
reported biological invasions from developing
nations is the result of insufficient research efforts
and inadequate data availability.
Over recent decades, many developing countries
have become potential sources and recipients of
invasive alien species to and from other countries of
the world due to fast economic globalisation of the
world and the associated acceleration in trade, travel,
and transport. However, for the majority of these
countries, even a preliminary inventory of alien
species does not yet exist (Wu et al. 2010). This is
even more crucial for the fast-emerging Asian
economies of China and India, which are at a higher
risk of biological invasions (Weber and Li 2008).
Over the last two decades, the expanding economy of
India has triggered large infrastructure development
projects that have led to the loss of natural habitats
and opening of disturbance corridors, providing
favourable habitats to the establishment and spread
of alien species (Sharma et al. 2005). Pertinently, a
recent report projected that India’s GDP per capita
will quadruple by 2020, and that the Indian GDP will
surpass that of the United States before 2050 (Podder
and Yi 2007).
As a step towards filling such knowledge gaps in
invasion biology research in developing countries,
including India, an inventory of alien species assumes
an urgent priority because it represents a critical
starting point for the scientific understanding and
systematic management of biological invasions at the
local, regional and global scales (Hulme et al. 2009a).
Such inventories, when compiled at the regional
scale, provide basic data necessary for testing various
scientific hypotheses in invasion biology (Cadotte
et al. 2006). These inventories, however, have to be
more than simple taxonomic checklists, and should
reflect the process of biological invasion (Palmer
et al. 1995; Pyšek et al. 2004). For this purpose, an
inventory needs to characterise the alien species at
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A. A. Khuroo et al.
different stages of the invasion process along the
introduction-naturalisation-invasion continuum (see
Richardson et al. 2000). Equally important is that
each alien species in an inventory is provided with
information about a suite of traits that captures the
species invasiveness and habitat invasibility. Further-
more, the inventory must be dynamic to allow
continuous updating. Once compiled, such an inven-
tory would provide an essential tool for early
detection and rapid response, informed decision-
making and sound policymaking in long-term mon-
itoring and systematic management of biological
invasions. As an example, the recently compiled
inventory of alien species of Europe under the project
DIASIE is the first continental-scale research effort
with far-reaching scientific, management and policy
implications (Hulme et al. 2009b).
While a number of research studies that inventorize
the alien flora have recently been carried out in China
(Liu et al. 2005, 2006; Weber et al. 2008; Wu et al.
2010) and in some other Asian countries (Corlett 1992;
Enomoto 1999; Koh et al. 2000; Wu et al. 2004), such
type of studies are lacking for India. Surprisingly, the
country’s Third National Report submitted to Con-
vention on Biological Diversity (CBD) states that 40%
of Indian flora is alien to the country, of which 21% is
invasive (CBD 2005). However, since a national alien
species inventory has not been compiled, the CBD
estimates are not based on reliable empirical data.
Prior to this Report, the estimates projected for the
proportion of alien plants in the Indian flora over the
last half a century have greatly varied (Chatterjee
1940; Maheshwari 1962; Nayar 1977; Saxena 1991).
In the absence of a baseline inventory, such estimates
are unreliable. This has serious implications for the
management of plant invasions because it hinders the
development of global indicators of biological inva-
sions (McGeoch et al. 2010). One of these indicators is
the number of invasive species in a region, which can
only be obtained by surveying and compiling alien
species inventories.
More recently, a few research studies on the alien
flora of some regions in India have been carried out
(Khuroo et al. 2007a, 2010; Negi and Hajra 2007;
Singh et al. 2010); and also, a preliminary list of
‘invasive alien flora of India’ has been compiled
(Reddy 2008). However, these regional studies are
narrow in scope and cannot be extrapolated for India
as a whole. The majority of these studies lacks an
Alien flora of India
explicit standard terminology and definition to char-
acterise the alien species at different stages of
invasion (see Pyšek et al. 2004), which limits their
applicability in comparative analyses, and in making
robust generalisations at the national and global
scales.
To fill up this knowledge gap, here we present a
comprehensive stage-based inventory of the alien
flora of India. The main advantage of such a stage-
based characterisation of alien flora is that it allows
the integration of species invasion status with the
relevant management option available (Khuroo et al.
2008). The specific objectives of the present study
were (a) to prepare a taxonomic conspectus of the
alien flora of India, (b) to characterise the alien
species with regard to their stage of invasion, and
(c) to explore biogeographic affiliations of the alien
flora of India.
Materials and methods
Study area
With an area of 3,287,240 sq km, India is the 2nd
largest country in Asia and 7th in the world, and has a
coastline of over 7,500 km length. It is the 2nd
largest populous country in the world, with human
population of more than 1 billion and population
density of 325 persons per sq km; 72% of the
population rural and 28% urban. The country is
administratively divided into 28 States and 7 Union
Territories (http://www.censusindia.gov.in).
India lies to the north of equator between 6° 450 –
37° 60 N latitude and 68° 70 –97° 250 E longitude. The
terrain of the country varies considerably, with
upland plains (the Deccan plateau) in the southern
region, flat to rolling plains along the Ganges river,
deserts in the west, and the Himalayan mountains in
the north. Notwithstanding regional climatic varia-
tions due to the geographical expanse and topograph-
ical heterogeneity, the major part of the country
experiences a tropical to sub-tropical climate. Three
distinct seasons are usually recognised in India: hot
summer (late April-late June), rainy monsoon (late
June-mid September), and mild winter (mid November-
late March). The country is situated at the conflu-
ence of three major terrestrial bio-realms, viz. the
Indo-Malayan, the Eurasian, and the Afro-tropical.
The vast geographical expanse of the country results
in a striking bio-climatic diversity, which ranges from
sea level to the highest mountain ranges in the world;
hot-arid conditions in the West to tropical evergreen
forests in North-east and Western Ghats; cold-arid
conditions in the trans-Himalayas to mangroves of
Sunderbans; and freshwater aquatic to marine eco-
systems (Negi 1986). Because of these factors, the
country shows a great habitat diversity represented by
forests, grasslands, wetlands, coastal, marine and
desert ecosystems harbouring a rich biodiversity.
India represents one of the world’s 12 Vavilovian
centres of origin and diversification of cultivated
plants, the ‘‘Hindustan Centre of origin of crop
plants’’ (Vavilov 1951); and has a representation of
12 biogeographical provinces and 5 biomes (Udvardy
1975). In fact, the country is one of the 17 megadi-
verse countries, ranking 3rd in Asia and 11th in the
world; and it shares four global biodiversity hotspots
(Sharma and Singh 2000; Mittermeier et al. 2005).
Since its independence in 1947, India has been the
world’s largest democracy; and presently it is one of
the fastest growing economies in the world. Despite
gains in economic development, the country faces
pressing problems, such as runaway overpopulation,
and environmental degradation. During the last few
decades, India has lost at least 50% of its pristine
forests, polluted over 70% of its water-bodies, and
converted much of its grasslands into agricultural
fields or urbanised townships (Sharma and Singh
2000). It is estimated that about 7.7% of the Indian
vascular-plant flora is facing risk of extinction (Rao
et al. 2003).
History of plant introductions
The entry of alien plants into India has occurred
through different pathways over time. A large
number of alien plants have migrated along ancient
and modern trade routes via land, sea and air, and
with general migration of pastoral and nomadic
tribes, with ballast, and as contaminants of food
grains, seeds of crop plants, etc. India’s colonial past
and historical trade relations have led to the plant
introductions by British, Portuguese, Spanish, French,
and from the Middle East and Central Asian countries
(Pandey 2000). During the sixteenth century, the
Portuguese had established a commercial route from
Lisbon to Brazil and thence from the Cape of Good
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Hope (South Africa) to Goa in India. They introduced
many tropical American plants, along with came a
number of aggressive weeds. Later on, the Spanish,
the Dutch, the French, and the British also brought
many economic plants from South America, Mexico
and Africa. There has also been intentional introduc-
tion of seeds and other propagules through botanic
gardens, arboreta, private seed companies, etc. Sea-
ports have always been a hub of alien plant
introduction, as ballast, ore, and coal piles, lumber-
yards and docks afford favourable conditions for
alien plants to establish in a new environment. The
unintentional introduction of invasive aliens has
resulted from contaminated garden seeds, food
grains, forage, and also by human transport systems
(Srivastava 1964).
Data sources
We developed this alien species inventory from an
extensive review of the literature (ca. 190 studies)
published over the last 120 years (1890–2010 AD).
Research papers published in peer-reviewed and local
journals, books, book chapters, scientific reports,
forest management plans, regional exotic floras, weed
floras, field guides, flower manuals, and other relevant
publications were reviewed and analysed (supple-
mentary material-I). The use of general Floras was
mostly avoided for two reasons. First, since the
publication of The Flora of British India (Hooker
1872–1897) more than a century ago, a complete and
updated Flora of India is still lacking. Second, until
recently, State and regional floras in the country rarely
indicate native or alien status of the species listed
therein. Very often, the naturalised alien species have
been treated as native to the floras. Therefore, we
focussed on the scientific literature dealing exclu-
sively with the alien flora. To characterise the
invasion status, the alien flora was annotated with
information extracted from the original sources and
additional literature such as research papers, local or
regional weed floras of different States in the country.
All species reported by earlier workers from different
regions of the country as being ‘exotic’, ‘non-indig-
enous’, ‘foreign’, ‘waifs’, ‘adventives’, ‘alien natura-
lised’, ‘introduced’, and ‘immigrants’ have been
included in our list of the alien plants of India. A
number of agricultural weed species, whose native
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A. A. Khuroo et al.
distribution range falls within the country have been
excluded from the inventory. Also, alien plants grown
exclusively under the green-house conditions, pots,
indoors, and other such artificial conditions have been
excluded from the inventory.
Only vascular plant species being alien to the
territory of the whole country (i.e. ‘Aliens to India’)
are included in the present study (see Pyšek et al.
2009). A large number of species recorded as aliens
in different regions of the country, but whose native
range falls within the political boundary of the
country (i.e. ‘Aliens in India’) have been excluded
in the present study. Example of an ‘Alien in India’
is the Himalayan Chir Pine (Pinus roxburghii)
recorded as ‘‘exotic’’ in southern India (Matthew
1969).
Taxonomy
The scientific nomenclature of species was updated
using taxonomic online databases, such as The Annual
Checklist of World Plants (http://www.sp2000.
org), International Plant Names Index (http://www.
ipni.org), E-Floras (http://www.efloras.org), and Germ-
plasm Resources Information Network (http://www.
grin.org). The taxa were considered at the taxonomic
rank of species; infra-specific taxa (subspecies, variety,
and forma, wherever recognised) have been subsumed
into their respective species, with the exception of a few
infra-specific taxa of Brassica oleracea. For the
arrangement of species within genera and of genera
within families, Mabberley (1997) has been followed.
However, the family Fabaceae (sensu lato) has been
divided into three families (sensu stricto): Papiliona-
ceae, Caesalpiniaceae and Mimosaceae.
Alien species recorded in the earlier literature with
invalid names, such as incorrect author citation,
misapplied name, etc., have been excluded from the
inventory, if later workers failed to validate these
species anywhere from the country. To ensure
reproducibility of the data, inclusion of name records
of alien species with conflicting taxonomy was
avoided. A high number of 520 scientific names,
previously recorded as distinct species, turned out to
be synonyms. This emphasizes the fact that if
synonymy is not taken into account, then it may lead
to taxonomic inflation in biodiversity studies (Khuroo
et al. 2007b).
Alien flora of India
Native ranges
In assigning the native ranges to all the species, a
biogeographic approach has been followed. Only
those species comprise the alien flora of India whose
native ranges fall outside the borders of the Indian
subcontinent. Instead of restricting to the narrow
political boundaries of India, the broad geographical
boundaries of the Indian subcontinent have been
taken into consideration. Therefore, the native spe-
cies from immediate neighbouring countries, such as
Pakistan, Nepal, Bhutan, and Bangladesh, were
excluded. Species native to one Indian region but
recorded as aliens in some other region(s) were
excluded from the present study. The native range for
each species was primarily obtained from the original
source of its record in India. However, in order to
minimise the error of judgement by earlier workers
about the alien status, and to cross-check the native
range records, the native ranges for all the species
were verified with data from the Germplasm
Resources Information Network (www.grin.org),
and some other recently published papers. After this
extensive exercise and critical review in light of
recent biogeographic evidences, a large pool of spe-
cies previously recorded as aliens to India turned out
to be native to the country and, therefore, were
excluded from the present study. Those alien species
where the recognition of native range was uncertain
were grouped under the cryptogenic category (Carlton
1996).
The records of native ranges for alien species are
of different quality in the literature. Historically, as
the species native ranges have been recorded at the
climatic, inter-continental, continental, sub-continen-
tal, country, provincial, regional, and sub-regional
scales, a single scheme of native ranges was imprac-
ticable for the entire alien flora. Above all, under
natural conditions, the species are distributed at
varying spatial scales, from one extreme of cosmo-
politan distribution to the other extreme of locally
restricted distribution. Therefore, the native ranges
for each species in the dataset were documented as
per the records available. However, for the biogeo-
graphic analysis of the total alien flora and the
invasive species separately, the native ranges were
pooled together at the continental scale (see Khuroo
et al. 2007a). In this scheme, the continent of South
America includes all the parts of tropical Central
America as well, and Asia includes all parts except
the Indian subcontinent.
Invasion status
We have compiled the dataset of all the alien plant
species in India, whether they grow only under
cultivation or have escaped into the wild. All the
alien species were designated as ‘aliens under
cultivation’ (Cl), unless there was any published
record or our own field observation of their escape
from cultivation, or self-reproducing population. The
remaining subset of alien species that escaped into
the wild has been evaluated with regard to their
invasion status in the country.
The terminology and definitions as proposed by
Pyšek et al. (2004) for designating the species as
casual (Cs), naturalised (Nt) and invasive (In) have
been followed. In addition, we defined two more
invasion status categories of casual or naturalised
(C/N) and naturalised or invasive (N/I), because for
many species information was not sufficient to
unambiguously allocate species to Cs, Nt, or In.
‘‘C/N’’ refers to those casual alien species for which
the current evidence is insufficient to be recognised
as naturalised but have the potential to become
naturalised in near future, and ‘‘N/I’’ refers to those
naturalised alien species for which the current
evidence is insufficient to be recognised as invasive,
but have the potential to become invasive in near
future. In summary, the present study uses six
different categories: cultivated (Cl), casual (Cs),
casual or naturalised (C/N), naturalised (Nt), natura-
lised or invasive (N/I), and invasive (In).
It is relevant to mention here that a large number
of alien plant species which in earlier studies have
been reported as casual, are reported as naturalised or
invasive in recent papers. For example, the species
reported as ‘occasional escapes’, or ‘run wild’ in
earlier studies have now become fully naturalised,
and many of these are spreading fast all over the
country. In such cases, the recent reports and field
observations have taken precedence in designation of
the invasion status. Many cultivated species, in
particular trees, that have long ‘established’ but have
no record of self-reproducing populations have been
called as ‘naturalised’ by earlier workers. All such
alien plant species in the present dataset, however,
have been put under the cultivated category (Cl), and
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 A. A. Khuroo et al.

excluded from naturalised category (Nt) (see Rich-
ardson et al. 2000).
To compare the intensity of plant invasions in
India with those of other countries of the world, we
calculated the species density as D = N/log (A);
where N is the total number of species at different
stages of invasion (casual to invasive), and A is the
area of the country (Rejmánek and Randall 1994).
Results
Taxonomic composition
The inventory includes 1,599 alien species in India;
these belong to 842 genera in 161 families (see
supplementary material-II). All the species are listed
alphabetically in their respective genera. For each
species is given its scientific name, author citation,
family, invasion status, and native range. (An MS-
excel file of the species list is available from the
corresponding author upon request). The angiosperms
are represented by 1,552 species belonging to 825
genera in 152 families; gymnosperms by 46 species
belonging to 16 genera in 8 families; and whereas
pteridophytes have a single species.
The three most species-rich families are Asteraceae
(134 spp.), Papilionaceae (114 spp.), and Poaceae (106
spp.). Of the total 161 families represented in this
inventory, the 20 largest families disproportionately
contribute 959 species, thereby showing a higher
average family:species ratio of 1:48 (Table 1). For the
remaining 141 families, sharing 640 species, the ratio
of family:species is very low (1:4.5). The first three
species-rich genera are Eucalyptus (25 spp.), Ipomoea
(22 spp.), and Senna (21 spp.). Of the total 842 genera
represented in the inventory, the largest 15 genera
disproportionately contribute 215 species, thereby
showing a higher average genus:species ratio of
1:14.3 (Fig. 1). For the remaining 827 genera, sharing
1,384 species, the genus:species ratio is very low
(1:1.7). Monospecific taxa, (i.e. taxa represented by a
single species in the inventory) are 546 genera and 49
families, which contribute 65% and 30% of the total

Table 1 Distribution of different invasion status categories, including cultivated, in the largest 20 families in the alien flora of India
Family Number of species
Total Cultivated Casual Casual or Naturalised Naturalised or Invasive
naturalised invasive

Asteraceae 134 25 9 6 25 26 43
Papilionaceae 114 48 3 5 38 9 11
Poaceae 106 19 15 10 34 15 13
Caesalpiniaceae 64 45 0 4 2 4 9
Solanaceae 63 24 3 4 12 7 13
Myrtaceae 59 51 0 3 5 0 0
Euphorbiaceae 52 23 1 3 7 4 14
Rubiaceae 42 30 0 0 5 5 2
Malvaceae 40 19 1 1 6 7 6
Apocynaceae 33 25 0 6 1 0 1
Convolvulaceae 30 7 1 1 11 0 10
Brassicaceae 29 9 6 0 7 2 5
Verbenaceae 29 14 1 9 1 1 3
Bignoniaceae 29 26 1 0 2 0 0
Amaranthaceae 28 2 1 3 3 5 14
Arecaceae 28 24 0 1 2 0 1
Rosaceae 23 16 0 5 2 0 0
Scrophulariaceae 20 9 1 1 4 0 5
Caryophyllaceae 18 4 4 0 3 4 3
Cupressaceae 18 15 0 3 0 0 0

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Alien flora of India

Fig. 1 Number of species Trifloium 9 (0)

at different stages of Cultivated
Brassica 9 (0) Casual
invasion, including Casual or Naturalised
Alternanthera 9 (7)
cultivated species, in the Naturalised
largest 15 genera in the
Oxalis 10 (5) Naturalised or invasive
Pinus 10 (0) Invasive
alien flora of India (total

Name of genera
Passiflora 10 (1)
number of alien species,
and number of invasive Bauhinia 10 (0)

species is given in Aristolochia 10 (0)

parentheses) Hibiscus 15 (0)

Euphorbia 16 (8)

Acacia 19 (2)

Solanum 20 (5)

Senna 21 (5)

Ipomoea 22 (7)

Eucalyptus 25 (0)

0 5 10 15 20 25
Number of species

Fig. 2 Numbers and Invasive

percentages of species at 225
Naturalised/ 14%
different stages of invasion, Invasive
including cultivated species, 134
Cultivated
in the alien flora of India 8%
812
51%

Naturalised
257
16%

Casual/
Naturalised
114 Casual
7% 57
4%

number of genera and families in the inventory,
respectively.
Invasion status
Out of the total alien species in the inventory, more
than half (812 spp., 51%) belong to the cultivated
category, i.e. having no report of their escape from
the cultivation. The distribution of the remaining 787
(49%) alien species under different stages of inva-
sion, with their species numbers and percentage in the
total alien flora, is as follows: casual (57 spp., 4%),
casual or naturalised (114 spp., 7%), naturalised (257
spp., 16%), naturalised or invasive (134 spp., 8%),
and invasive (225 spp., 14%) (Fig. 2).
There is a clear difference in the number and
percentage of species belonging to different families
and genera at various stages of invasion. The
proportion of species belonging to different invasion
categories shows great variation among the larger
families. While analysing contribution of the first 20
larger families to the Indian alien flora at different
stages of invasion in terms of number of species,
Myrtaceae has largest number of cultivated species
(51spp.), followed by Papilionaceae (48 spp.), and
Caesalpiniaceae (45 spp.). In contrast, Poaceae has

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 A. A. Khuroo et al.

Cultivated Casual Casual or Naturalised Naturalised Naturalised or Invasive Invasive

Rosaceae 0.00

Cupressaceae 0.00
Bignoniaceae 0.00
Myrtaceae 0.00
Apocynaceae 3.03
Arecaceae 3.57
Rubiaceae 4.76

Papilionaceae 9.65
Verbenaceae 10.34
Poaceae 12.26

Caesalpiniaceae 14.06

Malvaceae 15.00
Brassicaceae 17.24
Caryophyllaceae 16.67
20.63
Solanaceae
25.00
Scrophulariaceae
Euphorbiaceae 26.92
Asteraceae 32.09

Convolvulaceae 33.33
Amaranthaceae 50.00

0% 20% 40% 60% 80% 100%

Fig. 3 Percentage contribution of the first 20 larger families to the different invasion status categories, including cultivated, in the
alien flora of India (percentage of invasive species in the families is given on the right hand side)

the largest number of casual species (15 spp.),
followed by Asteraceae (9 spp.), and Brassicaceae
(6 spp.). In the case of casual or naturalised category,
again Poaceae has the largest number of 10 species,
followed by Verbenaceae (9 spp.), and Asteraceae
and Apocyanceae (6 spp. each). In the case of
naturalised category, Papilionaceae has the largest
number of 38 species, followed by Poaceae (34 spp.),
and Asteraceae (25 spp.). In the case of naturalised or
invasive category, Asteraceae has the largest number
of 26 species, followed by Poaceae (15 spp.), and
Papilionaceae (9 spp.). Finally, in the case of invasive
category, again Asteraceae has the largest number of
43 species, followed by Amaranthaceae and Euphor-
biaceae (14 spp. each), and Poaceae and Solanaceae
(13 spp. each) (Table 1).
While as Asteraceae contributes the largest num-
ber of species (43 spp.) to the list of 225 invasive
species, it ranks third (32%) much after Convolvul-
aceae (33%) and Amaranthaceae (50%), in terms of
proportion of invasive species to the total number of
species in the alien flora. Similarly, while Papilion-
aceae and Poaceae rank third in terms of species
numbers, these families have rather few invasive
species and rank distant eleventh and twelfth in their
percentage contribution to the invasive species. There
are eight families having more than 70% of cultivated
species (Fig. 3). Whereas Eucalyptus (25 spp.) is the
largest genus in the alien flora, it does not contribute
even a single invasive species. On the other hand,
Euphorbia (16 spp.) contributes 8 invasive species.
Some other species-rich genera, such as Alternan-
thera (9 spp.) and Ipomoea (22 spp.) have 7 invasive
species each; and Oxalis (10 spp.), Senna (21 spp.)
and Solanum (20 spp.) have 5 invasive species each
(Fig. 1).
In the alien flora of India, 49 families are
represented by a single species, the majority of
which (28 spp.) belongs to the cultivated category.
The remaining 21 species are distributed among
different stages of invasion (Fig. 4a). Interestingly,
some of these families comprise only invasive species
in India (e.g., Balsaminaceae, Impatiens balsamina
L.; Cannabaceae, Cannabis sativa L.; Ceratophylla-
ceae, Ceratophyllum demersum L.; Martyniaceae,
Martynia annua L.; Menispermaceae, Cissampelos
pareira L.; Salvinaceae, Salvinia molesta Mitchell;
and Potamogetonaceae, Potamogeton crispus L.).

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Alien flora of India

Fig. 4 Number of 30 28
monospecific families a by b
their invasion status and
a
25

Number of monotypic taxa

b by their native region
(continental) in the alien
20
flora of India 20 19

15

10
7
6
5 5
5 3
2 2
1
0
0

Casual

Africa

Asia
Casual or naturalised

Europe
Cultivated

Australia

North America

South America
Invasive
Naturalised or Invasive
Naturalised

Invasion status Native region (Continental)

In terms of percentage of alien species to the total Schinus terebinthifolius, Spathodea campanulata) are
number of species in India, only two families, among the cultivated species in India.
Martyniaceae and Turneraceae show a value of
100%; and six families (Caesalpiniaceae, Oxalida- Biogeographic affiliations
ceae, Pontederiaceae, Portulacaceae, Solanaceae,
Typhaceae) show a value of more than 50%. However, More than one-third (35%) of the total alien flora of
when analysed in terms of percentage of invasive India has its origin in the South American continent.
species to the total number of alien species in India, Most South American species come from Mexico and
seven families have a value of 100%, e.g. Balsamin- Brazil (75 and 54 species, respectively). The per-
aceae, Cannabaceae, Martyniaceae, Pontederiaceae, centage contribution by other continents greatly
Potamogetonaceae, Salvinaceae and Tureraceae. varies (Fig. 5); other important source areas are Asia
Three families (Asclepiadaceae, Capparaceae and and Africa. Cryptogenic species make 1%. More
Oxalidaceae) show more than 50% values (Table 2). specifically, while analysing the native ranges of 225
Out of the 36 plant species recognised among the invasive species, more than half (52%) are from
‘‘world’s worst invasive alien species’’ (Lowe et al. South America. The contribution of other continents
2000), the present inventory of alien vascular plant is as follows: Africa (16%), Asia (16%), Europe
species in India includes 17 species. Of these, the (9%), North America (4%), Australia (2%) and
present study characterized 11 species as invasive to cryptogenic (1%) (Fig. 5). The pattern of origins is
India: Acacia mearnsii, Arundo donax, Chromolaena very similar if only those families represented by
odorata, Clidemia hirta, Imperata cylindrica, Lan- single species in the Indian alien flora are considered
tana camara, Leucaena latisiliqua (syn. Leucaena (Fig. 4b).
leucocephala), Mikania micrantha, Opuntia stricta,
Ulex europeus and Sphagneticola trilobata (syn.
Wedelia trilobata); of the remaining 2 species— Discussion
Mimosa pigra and Prosopis glandulosa, are recogni-
sed as naturalised or invasive, whereas the four other The 1,599 alien species recorded in this study lead to
species (Cinchona pubescens, Psidium cattleyanum, a proportion of 8.5% alien species in the vascular

123
 A. A. Khuroo et al.

Table 2 Numerical and proportion values of families with representation of invasive species in the alien flora of India, with
reference to the total number of alien species, and total number of species in India
Family Total number of Total number Proportion of alien species Total number of Proportion of
species in India of alien species to total number of species invasive species invasive to alien
species

Acanthaceae 500 16 3.20 1 6.25

Amaranthaceae 60 28 46.67 14 50.00
Apiaceae 288 10 3.47 2 20.00
Apocynaceae 119 33 27.73 1 3.03
Araceae 126 9 7.14 1 11.11
Arecaceae 94 28 29.79 1 3.57
Asclepiadaceae 260 5 1.92 4 80.00
Asteraceae 800 134 16.75 43 32.09
Balsaminaceae 200 1 0.50 1 100.00
Boraginaceae 209 13 6.22 1 7.69
Brassicaceae 207 29 14.01 5 17.24
Cactaceae NA 13 NA 2 15.38
Caesalpiniaceae 92 64 69.57 9 14.06
Cannabaceae NA 1 NA 1 100.00
Capparaceae 55 5 9.09 4 80.00
Caryophyllaceae 122 18 14.75 3 16.67
Casuarinaceae 12 3 25.00 1 33.33
Ceratophyllaceae 2 1 50.00 1 100.00
Chenopodiaceae 71 4 5.63 2 50.00
Convolvulaceae 199 30 15.08 10 33.33
Cyperaceae 1,545 15 0.97 3 20.00
Euphorbiaceae 527 52 9.87 14 26.92
Lamiaceae 435 16 3.68 3 18.75
Liliaceae 214 10 4.67 1 10.00
Malvaceae 93 40 43.01 6 15.00
Martyniaceae 1 1 100.00 1 100.00
Melastomataceae 150 3 2.00 1 33.33
Menispermaceae 43 1 2.33 1 100.00
Mimosaceae 127 44 34.65 7 15.91
Moraceae 122 9 7.38 1 11.11
Nyctaginaceae 15 5 33.33 1 20.00
Oxalidaceae 19 11 57.89 6 54.55
Papaveraceae 27 7 25.93 2 28.57
Papilionaceae 973 114 11.72 11 9.65
Passifloraceae 24 10 41.67 1 10.00
Pedaliaceae 4 2 50.00 1 50.00
Piperaceae 101 3 2.97 1 33.33
Poaceae 1,291 106 8.21 13 12.26
Polygonaceae 164 12 7.32 1 8.33
Pontederiaceae 3 2 66.67 2 100.00
Portulacaceae 8 5 62.50 2 40.00
Potamogetonaceae 18 1 5.56 1 100.00

123
Alien flora of India

Table 2 continued
Family Total number of Total number Proportion of alien species Total number of Proportion of
species in India of alien species to total number of species invasive species invasive to alien
species

Primulaceae 183 2 1.09 1 50.00

Rubiaceae 616 42 6.82 2 4.76
Salvinaceae NA 1 NA 1 100.00
Sapindaceae 55 11 20.00 1 9.09
Scrophulariaceae 368 20 5.43 5 25.00
Simaroubaceae 16 3 18.75 1 33.33
Solanaceae 75 63 84.00 13 20.63
Tiliaceae 53 9 16.98 6 66.67
Turneraceae 2 2 100.00 2 100.00
Typhaceae 3 2 66.67 1 50.00
Urticaceae 153 3 1.96 1 33.33
Verbenaceae 140 29 20.71 3 10.34
Zygophyllaceae 15 5 33.33 1 20.00
NA not available

60
Total alien flora 52
Invasive species only
50
Proportion (%)

40 35

30

21
20

20 16 16

11
8 9
10
4
2 4
1 1

0
Asia Australia Africa Europe North America South America Cryptogenic

Continent

Fig. 5 Proportion of total alien species and only invasive species from different continents (including cryptogenic) contributing to
the alien flora of India

flora of India, based on a total number of 18,748
vascular plant species (Arora and Bhatt 2008). The
percentage is only 4.2% when calculated for the alien
flora excluding cultivated species (i.e. total number of
species from casual to invasive status is 787 spp.) to
the total vascular flora of India; and 14% (225 spp.)
of the total alien flora having reached the invasive
stage (Fig. 2). These percentages are significantly
lower than previously published estimates. For
instance, according to the Third National Report,
40% of the Indian flora is alien, out of which 21% is
invasive. That would lead to ca.7,500 alien species in

123

India, out of which ca. 1,575 would be invasive
species. These estimates, however, are more likely
based on assumptions without sound empirical data.
The percentage of 4.2% calculated for the alien
flora of India, excluding cultivated species, is much
higher than that reported for China (1.4%) (percent-
age based on 420 species reported in Wu et al.
(2010), and 31,000 vascular plant species recorded in
Flora of China) (http://hua.huh.harvard.edu/china/).
The total number of 1,599 species alien in India is
lower than the 28,866 species recorded as alien in
Australia (Randall 2007), 25,049 species in New
Zealand (Diez et al. 2009), and 2,843 species in
Europe (Lambdon et al. 2008). Such a lower number
and percentage of alien species in the Indian flora
compared to numbers from the developed world may
possibly be due to: (1) non-inclusion of archaeo-
phytes in the present study, (2) poor record-keeping
of alien species introduced into the country by vari-
ous governmental and private agencies, (3) the
records of introduced plants having been traditionally
least-reported and/or excluded from the floristic
works, and (4) very scarce taxonomic research effort
in developing countries, including India.
In concordance with the alien floras of Europe
(Lambdon et al. 2008) and China (Wu et al. 2010),
Asteraceae is the most species-rich family in the alien
flora of India. However, Papilionaceae is the second
species-rich family, in contrast to Poaceae in Europe
and China, which ranks third in India. Brassicaceae,
which is the fifth largest family in Europe and the
sixth in China, does not appear among the top ten
largest families in India. Rosaceae, which is the third
largest family in the European alien flora, has rather
few species (rank 17) in India. Brassicaceae and
Rosaceae are the families mostly concentrated in the
northern temperate hemisphere and their lesser rep-
resentation in India compared to Europe may possibly
be due to only few montane areas in northern India,
having influence of temperate climate.
It is a well known fact that the higher species
contribution of families such as Asteraceae, Papi-
lionaceae or Poaceae to the alien floras is mainly due
to a sampling effect. These families are globally
known to be species-rich and, therefore, higher
number of alien species belonging to these families
is expected. However, the percentage contribution of
the taxonomic families to the invasive species is
disproportionate. Thus, half of the species (50%) in
123
A. A. Khuroo et al.
the family Amaranthaceae are invasive, followed by
nearly one-third species in Convolvulacaeae (33%),
and Euphorbiacaeae (27%). Such a higher predispo-
sition of species from Amaranthaceae, Convolv-
ulacaeae and Euphorbiaceae to be invasive than
others has also been reported from China (Wu et al.
2010). On the other hand, Papilionaceae and Poaceae,
ranking 2nd and 3rd in terms of species number,
show a lesser percentage contribution of 10 and 12%
to the invasive species, respectively.
Amongst the highly represented genera in the alien
flora of India, only Euphorbia, Solanum and Trifo-
lium are reported in Europe; while in China,
predominance of genera such as Euphorbia, Senna,
Alternanthera, Ipomoea has been reported. However,
Oenothera and Amaranthus—predominant genera in
the alien flora of Europe and China—are interesting
exceptions, as these are not amongst the species-rich
genera in the alien flora of India (Fig. 1). Taxonomic
distribution of nearly 60% of total species in the first
20 largest families indicates a phylogenetic clustering
in the alien flora of India (Table 1). Whether this is
due to human preference for the introduction of
particular type of taxa, or something else, merits
detailed investigation, which is beyond the scope of
the present study.
Our study is significant in the sense that it includes
the alien cultivated plant species, which have often
been ignored in most of studies on alien floras. It is
this species pool of cultivated plants which is the
potential source of future invasive species. In
essence, many alien species presently escaped from
cultivation and recognised at different stages of inva-
sion were under cultivation at some point of time.
Comparative analyses of failure and success of spe-
cies to escape from cultivation is possible only with
the inclusion of ‘species under cultivation’ in the
alien flora inventories (Diez et al. 2009). Moreover,
rigorous testing of hypotheses and robust scientific
understanding in invasion biology can be achieved
only when the data is made available for the entire
pool of alien species, their residence time, and extent
of naturalisation.
Among the 36 plant species belonging to the
‘‘world’s worst invasive alien species’’, 17 species
also occur in India. Of these, the recognition of 11
species as invasive and two species as naturalised or
invasive is a considerably higher number. Immedi-
ately relevant from the management perspective is
Alien flora of India
that the four alien species Cinchona pubescens,
Psidium cattleyanum, Schinus terebinthifolius and
Spathodea campanulata, which are not reported to
have escaped from cultivation in India, need to be
monitored strictly and at best, their cultivation should
be discouraged as a preventive measure.
The total number of naturalised species has been
used as a reliable predictor of invasive species.
Rejmánek and Randall (2004) concluded that
15–30% of the total naturalised species were invasive
in the US, a proportion higher than the 10% proposed
by Williamson and Fitter (1996). Our results are in
concordance with those of Rejmánek and Randall
(2004), as 31% (225 spp.) of the total naturalised
species (730 spp.) belonged to the invasive category,
when the 869 species (cultivated 812 spp., casual 57
spp.) were excluded from the analysis.
On analysing the intensity of plant invasions, using
the density index (Rejmánek and Randall 1994),
India shows a lower value of 120.76 as compared to
298.95 for New Zealand, 197.13 for Japan, and
175.36 for British Isles (Wu et al. 2004). On the other
side, the density value of 120.76 for India is much
higher than that for China (60.14), (Wu et al. 2010).
Such a higher value may be the result of relatively
longer history of colonisation by the Europeans, and
rulers from the Central Asia in India than in China. In
addition, until the recent past, China was much more
isolated from rest of the world than India.
By and large, the Indian alien flora has a South
American origin; though the floristic elements of
Asian, African, European, Australian, and North
American origin are also well represented in the
total alien flora, as well as in the list of invasive
species. Nevertheless, the proportion of South Amer-
ican species, in particular tropical Central America, is
disproportionately higher in the case of invasive
species (52%), as compared to the total alien flora
(35%). Opposite to this trend, the proportion of
Australian species is disproportionately higher in the
case of total alien flora (8%), as compared to invasive
species (2%). The possible explanation for the
maximum proportion of species from South America
can be (1) the higher propagule pressure from
different countries, such as Brazil and Mexico, to
India via historical trade routes through the human
agency of European colonisers and traders, and (2)
more or less matching of similar tropical climate.
When considering that 56% of species originate from
the Americas, the results in the present study are
similar to those reported for China, wherein 58% of
species have their origin from the Americas (Wu
et al. 2010). On the other hand, however, the share of
American species in the alien flora of Europe is
34.8% (Lambdon et al. 2008), much lower than in the
present study. Such a varying trend can be explained
in terms of larger influence of tropical climate in
India and China, rather than in Europe.
Future implications
The inventory of the alien flora of India will help in
bridging the geographical knowledge gaps in inva-
sion biology research. The inventory will serve as a
scientific baseline for investigating the patterns,
pathways, extent, impacts and effective management
of plant invasions in India. As the inventory is based
on the synthesis of knowledge accumulated over the
last one century, it will stimulate the much-needed
research on invasion biology, invite attention of
policymakers and raise public awareness in the
developing world, including India. The characteriza-
tion of alien flora at different stages of invasion, as
attempted in the present study, will help in targeted
prioritisation of research and management efforts
required at the country scale. It will also pave way for
formulation of a predictive management framework
that includes elements of risk analysis, early detection
and warning systems which are pre-requisites for
taking informed decisions for the eradication and
control of invasive species. Finally, the availability of
data on invasive alien plant species in India will
contribute to the larger goal of setting robust Global
Biodiversity Targets and Indicators, with far-reaching
policy implications for the conservation and sustain-
able use of biodiversity.
Acknowledgments We are highly thankful to Prof. Marcel
Rejmánek, University of California, Davis, for his valuable
comments on the earlier version of the manuscript. Dr. Marc
Cadotte, Associate Editor and two anonymous reviewers
deserve thanks for their useful suggestions and critical
comments which greatly improved the manuscript.
Taxonomic expertise shared by Dr. C. S. Reddy, NRSA,
Hyderabad (India) is greatly acknowledged. The Head,
Department of Botany, University of Kashmir, J & K, India,
is acknowledged for providing the necessary facilities during
the course of present study. Lastly, AAK acknowledges the
kind help, valuable suggestions and research papers provided
123

by the researchers and staff working in different institutions,
herbaria and libraries of the country, and abroad.
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